长期水分胁迫对小麦生育中后期根叶渗透调节能力、渗透调节物质的影响

以 2个抗旱性强的和 2个抗旱性弱的小麦品种为材料 ,研究了中度及严重水分胁迫对根系及叶片渗透调节能力的影响。结果表明 :随着水分胁迫的加剧 ,叶片的渗透调节能力增强 ,但在籽粒迅速扩大的灌浆期 ,叶片的渗透调节能力下降。去穗处理明显地提高叶片的渗透调节能力。说明叶片渗透调节能力的高低与同化物的供应及分配有关。不同品种根系渗透调节能力与叶片基本一致 ,但根系的渗透调节能力低于叶片。开花、灌浆期根系的渗透调节能力大大降低 ,严重水分胁迫下根系的渗透调节能力低于中度水分胁迫。这一方面与同化物的供应有关 ,另一方面严重水分胁迫还会对根细胞造成损伤 ,对根系的渗透调节能力产生影响。渗透调节物质的变化趋势与渗透调节能力基本一致。叶片中 K+对渗透调节的贡献最大 ;其次是可溶性糖 ,6种渗透调节物质排列顺序为 K+>可溶性糖 >游离氨基酸 >Ca2 +>Mg2 +>Pro。根系中仍以 K+占绝大部分 ,但根系中 Ca2 +也是不可忽视的成分之一。     

The cooperative relation between non-hydraulic root signals and osmotic adjustment under water stress improves grain formation for spring wheat varieties

Non-hydraulic root signals (nHRS) and osmotic adjustment (OA) are two important adaptive responses of plants to water stress. There is little understanding of their relationships during water stress. The threshold range of soil water potential to occurrence of nHRS, the capacity for OA, grain yield and water use efficiency (WUE) were examined in three spring wheat ( Triticum aestivum L.) varieties (two bred after 1975 and one bred before 1900) under water stress conditions. The threshold range of nHRS was significantly correlated with the maintenance rate of grain yield (MRGY) ( r  = 0.99, P  < 0.05) under moderate drought (−0.49 to −0.55 MPa) but not under severe drought (−0.70 to −0.76 MPa). There were similar correlations between OA and the MRGY. However, regulation of nHRS precedes OA during gradual water stress. The threshold range of nHRS and OA was positively correlated ( r  = 0.93, P  < 0.05), suggesting a mechanism for adapting to drought. WUE was higher for modern than for old varieties and was correlated with the root efficiency (full biomass weight including root per root weight, r  = 0.78, P  < 0.05) and the root water uptake efficiency (water consumption per root weight, r  = 0.72, P  < 0.05). However, there was a significant negative correlation between WUE and root weight ( r  = –0.84, P  < 0.01). The cooperative relationship between the threshold range of nHRS and OA under water stress was beneficial for improving grain formation for spring wheat varieties.     

The possible role of hydroxylation in the detoxification of atrazine in mature vetiver (Chrysopogon zizanioides Nash) grown in hydroponics

The resistance mechanism of vetiver (Chrysopogon zizanioides) to atrazine was investigated to evaluate its potential for phytoremediation of environment contaminated with the herbicide. Plants known to metabolise atrazine rely on hydroxylation mediated by benzoxazinones, conjugation catalyzed by glutathione-S-transferases and dealkylation probably mediated by cytochromes P450. All three possibilities were explored in mature vetiver grown in hydroponics during this research project. Here we report on the chemical role of benzoxazinones in the transformation of atrazine. Fresh vetiver roots and leaves were cut to extract and study their content in benzoxazinones known to hydroxylate atrazine, such as 2,4-dihydroxy-2H-1,4-benzoxazin-3(4H)-one (DIBOA), 2,4-dihydroxy-7-methoxy-2H-1,4-benzoxazin-3(4H)-one (DIMBOA) and their mono- and di-glucosylated forms. Identification of benzoxazinones was performed by thin layer chromatography (TLC) and comparison of retention factors (Rf) and UV spectra with standards: although some products exhibited the same Rf as standards, UV spectra were different. Furthermore, in vitro hydroxylation of atrazine could not be detected in the presence of vetiver extracts. Finally, vetiver organs exposed to [14C]-atrazine did not produce any significant amount of hydroxylated products, such as hydroxyatrazine (HATR), hydroxy-deethylatrazine (HDEA), and hydroxy-deisopropylatrazine (HDIA). Altogether, these metabolic features suggest that hydroxylation was not a major metabolic pathway of atrazine in vetiver.     

Different mechanisms of adaptation to cyclic water stress in two South Australian bread wheat cultivars

In the South Australian wheat belt, cyclic drought is a frequent event represented by intermittent periods of rainfall which can occur around anthesis and post-anthesis in wheat. Three South Australian bread wheat (Triticum aestivum L.) cultivars, Excalibur, Kukri, and RAC875, were evaluated in one greenhouse and two growth-room experiments. In the first growth-room experiment, where plants were subjected to severe cyclic water-limiting conditions, RAC875 and Excalibur (drought-tolerant) showed significantly higher grain yield under cyclic water availability compared to Kukri (drought-susceptible), producing 44% and 18% more grain compared to Kukri, respectively. In the second growth-room experiment, where plants were subjected to a milder drought stress, the differences between cultivars were less pronounced, with only RAC875 showing significantly higher grain yield under the cyclic water treatment. Grain number per spike and the percentage of aborted tillers were the major components that affected yield under cyclic water stress. Excalibur and RAC875 adopted different morpho-physiological traits and mechanisms to reduce water stress. Excalibur was most responsive to cyclic water availability and showed the highest level of osmotic adjustment (OA), high stomatal conductance, lowest ABA content, and rapid recovery from stress under cyclic water stress. RAC875 was more conservative and restrained, with moderate OA, high leaf waxiness, high chlorophyll content, and slower recovery from stress. Within this germplasm, the capacity for osmotic adjustment was the main physiological attribute associated with tolerance under cyclic water stress which enabled plants to recover from water deficit.     

低水势下冬小麦胚芽鞘长度与叶片的渗透调节能力及大田产量关系的研究

本文以７个抗旱性不同的冬小麦品种（系）为材料,研究了低水势下芽鞘长度与大田干旱条件下叶片的渗透调节能力及大田产量的关系。结果表明：低水势下芽鞘长度与叶片的渗透调节能力及产量有极为密切的关系。为冬小麦抗旱鉴定及抗旱个体筛选提供了一个良好的指标。     

Benzoxazinoids-cyclic hydroxamic acids, lactams and their corresponding glucosides in the genus Aphelandra (Acanthaceae)

An improved method of sample preparation and simultaneous HPLC separation was developed that allowed the separation of 2,4-dihydroxy-1,4-benzoxazine-3(4H)-one (DIBOA), 2,4-dihydroxy-7-methoxy-1,4-benzoxazine-3(4H)-one (DIMBOA), 2-hydroxy-1,4-benzoxazine-3(2H)-one (HBOA), 2-hydroxy-7-methoxy-1,4-benzoxazine-3(2H)-one (HMBOA) and their corresponding glucosides as well as the benzoxazolinones BOA and MBOA. The amount and distribution of these compounds was determined in the roots of Aphelandra squarrosa and A. fuscopunctata plants. There is a significant difference in the amount and distribution of this substance class in the two species analyzed. The results are discussed in relation to their function as defence compounds and allelochemicals.     

<Emphasis Type="Italic">ScBx</Emphasis> gene based association analysis of hydroxamate content in rye (<Emphasis Type="Italic">Secale cereale</Emphasis> L.)

Hydroxamates (HX) are major secondary metabolites synthesized by rye and are responsible for some of the unique properties of this cereal, including good tolerance of biotic and abiotic stresses and allelopathy. Recently, five genes encoding enzymes taking part in HX biosynthesis have been sequenced and characterized, which was the starting point to undertake the present study. Association analysis of the content of six HX–HBOA (2-hydroxy-1,4-benzoxazin-3-one), GDIBOA (2,4-dihydroxy-1,4-benzoxazin-3(4H)-one glucoside), DIBOA (2,4-dihydroxy-1,4-benzoxazin-3(4H)-one), GDIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3(4H)-one glucoside), DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3(4H)-one) and MBOA (6-methoxy-benzoxazolin-2(3H)-one) in the above-ground parts of plants and roots was performed on a population consisting of 102 and 121 diverse inbred lines, in 2013 and 2014, respectively. Altogether, 48 single nucleotide polymorphisms (SNPs) were found to be associated with the content of at least one HX: 20 SNPs were associated with HX synthesized in the above-ground parts of rye plants (AG-SNP), and 28 were associated with HX synthesized in the roots (R-SNP). The highest number of SNPs was present in genes ScBx1 (9) and ScBx5 (11). The majority of SNPs were affected by environmental factors, except for two: ScBx4_1702 associated with GDIBOA and MBOA contents, and ScBx5_1105 associated with HBOA content in roots.     

Osmotic adjustment is a prime drought stress adaptive engine in support of plant production

Osmotic adjustment (OA) and cellular compatible solute accumulation are widely recognized to have a role in plant adaptation to dehydration mainly through turgor maintenance and the protection of specific cellular functions by defined solutes. At the same time, there has been an ongoing trickle of skepticism in the literature about the role of OA in supporting crop yield under drought stress. Contrarian reviews argued that OA did not sustain turgor or that it served mainly for plant survival rather than productivity. This critical review examined 26 published studies where OA was compared with yield under drought stress in variable genotypes of 12 crops, namely, barley, wheat, maize, sorghum, chickpea, pea, pigeon pea, soybean, canola, mustard, castor bean and sunflower. Over all crops a positive and significant association between OA and yield under drought stress were found in 24 out of 26 cases. Considering that it is generally difficult to find a singular plant trait responsible for yield advantage of numerous crops under different drought stress conditions, this evidence is no less than remarkable as proof that OA sustains crop yield under drought stress.     

Improvement in drought tolerance in bread wheat is related to an improvement in osmolyte production,antioxidant enzyme activities,and gaseous exchange

Water deficit stress negatively affects wheat growth, physiology, and yield. In lab and hydroponic experiments, osmotic stress levels (control, −2, −4, −6 and −8 Bars) created by PEG-6000, caused a significant decline in germination, mean germination time, root, shoot, and coleoptile length in both wheat genotypes examined. Germination was inhibited more in Wafaq-2001 than in Chakwal-50. Wafaq-2001 showed a higher susceptibility index based on root and shoot dry weight than did Chakwal-50. Wheat plants exhibited osmotic adjustment through the accumulation of proline, soluble sugars, soluble proteins, and free amino acids, and increased antioxidation activities of superoxide dismutase, peroxidase, catalase, and malondialdehyde. Increasing water deficit stress caused a linear decline in chlorophyll contents, leaf membrane stability, and relative water content in all wheat plants, with Wafaq-2001 showing a more severe negative impact on these parameters with increasing stress levels. The results suggest the possibility of utilizing some of these parameters as quantitative indicators of water stress tolerance in plants. Gas exchange measurements (photosynthesis, transpiration, stomatal conductance), leaf osmotic potential, water potential, and yield attributes decreased more abruptly with increasing water deficit, whereas leaf cuticular wax content increased in both genotypes, with more severe impacts on Wagaq-2001. More reduction in biochemical, physiological, and yield attributes was observed in Wafaq-2001 than was observed in Chakwal-50. Based on these results, we can conclude that Chakwal-50 is a more drought-tolerant genotype, and has excellent potential for future use in breeding programs to improve wheat drought tolerance.     

小麦叶片光合作用与其渗透调节能力的关系

在缓慢干旱条件下,小麦叶片渗透调节能力在一定范围内随胁迫程度的加剧而增加,而在快速干旱下,渗透调节能力丧失。小麦叶片通过渗透调节使光合速率和气孔导度对水分胁迫的敏感性降低,叶片维持较高的电子传递能力、RuBP羧化酶活性和叶绿体光合能量转换系统活性,并推迟了小麦叶片光合速率受气孔因素限制向叶肉细胞光合活性限制转变的时间。     

Leaf water characteristics and drought acclimation in sunflower genotypes

The responses of leaf water parameters to drought were examined using three sunflower (Helianthus annuus L.) genotypes. Osmotic potential at full water saturation (π¹⁰⁰), apoplastic water fraction (AWF) and bulk elastic modulus (BEM) were determined by pressure-volume curve analysis on well watered or on water-stressed plants (−1.0 MPa Ψ₁ < −1.5 MPa) previously drought-pretreated or not. The drought-pretreated plants were subjected to a 7-day drought period (predawn leaf water potential reached −0.9 MPa) followed by 8 days of rewatering. In well watered plants, all genotypes in response to drought acclimation displayed a significantly decreased π¹⁰⁰ associated with a decrease in the leaf water potential at the turgor-loss point (decrease in Ψ_tlp was between 0.15 and 0.21 MPa, depending on the genotype). In two genotypes, drought acclimation affected the partitioning of water between the apoplastic and symplastic fractions without any effect on the total amount of water in the leaves. As a third genotype displayed no modification of AWF and BEM after drought acclimation, the decreased π¹⁰⁰ was only due to the net accumulation of solutes and was consistent with the adjustment of the photochemical efficiency observed previously in this genotype in response to drought acclimation. In water-stressed plants, the osmotic adjustment (OA) can increase further beyond that observed in response to the drought pretreatment. However, the maintenance of photosynthetic rate and stomatal conductance at low leaf water potentials not only depends on the extent of osmotic adjustment, but also on the interaction between OA and AWF or BEM. Adaptative responses of leaf water parameters to drought are thus quite contrasted in sunflower genotypes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Occurrence and Characterization of 2-Hydroxy-1,4-benzoxazin-3-one and Indole Hydroxylases in Juvenile Wheat

Cyclic hydroxamic acids, 2,4-dihydroxy-1,4-benzoxazin-3-one (DIBOA) and its 7-methoxy analogue (DIMBOA), occur transiently in high amounts in wheat and maize during the juvenile, non-autotrophic stage of growth. To elucidate the biosynthetic enzymes operating for the transient production of these compounds, we examined the hydroxylating activities for 2-hydroxy-1,4-benzoxazin-3-one (HBOA), the immediate precursor of DIBOA, and indole, the first intermediate of the biosynthetic pathway that branches off from the tryptophan pathway, by using microsomes prepared from wheat seedlings. Both hydroxylases occurred soon after germination, reached a maximum 48 h after germination, and decreased to finally disappear as the plants grew into the autotrophic growth stage. The mode of appearance and disappearance similar to that of hydroxamic acids, suggesting that elevated expression of the whole set of enzymes involved in the biosynthesis after indole was responsible for the transient occurrence of hydroxamic acids. The hydroxylating activity was also observed for 1,4-benzoxazin-3-one, a putative precursor of HBOA, but to significantly less extent than that for HBOA and indole.     

Effect of light on DIMBOA-glucoside concentration in wheat {Triticum aestivum L.)

DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) glucoside in wheat has been suggested to play a role in plant resistance to cereal aphids. Thus there is considerable interest in trying to breed modern wheat varieties with increased concentrations of this compound. To further such work we assessed the effects of light intensity on DIMBOA glucoside production in wheat seedlings under both indoor and outdoor conditions. Light had a marked effect on DIMBOA-glucoside concentrations, with levels in plants being negatively related to light intensity. Our findings suggest that breeding aimed at increasing levels of DIMBOA-glucoside would be most effective if test plants for selection were to be grown under weak light conditions.     

Phenotyping shows improved physiological traits and seed yield of transgenic wheat plants expressing the alfalfa aldose reductase under permanent drought stress

Members of the aldo–keto reductase family including aldose reductases are involved in antioxidant defense by metabolizing a wide range of lipid peroxidation-derived cytotoxic compounds. Therefore, we produced transgenic wheat genotypes over-expressing the cDNA of alfalfa aldose reductase gene. These plants consequently exhibit 1.5–4.3 times higher detoxification activity for the aldehyde substrate. Permanent drought stress was generated in the greenhouse by growing wheat plants in soil with 20 % water capacity. The control and stressed plants were monitored by a semi automatic phenotyping platform providing computer-controlled watering, digital and thermal imaging. Calculation of biomass values was based on the correlation (R ² = 0.7556) between fresh weight and green pixel-based shoot surface area. The green biomass production by plants of the three transgenic lines was 12–26–41 % higher than the non-transgenic plants’ grown under water limitation. Thermal imaging of stressed non-transgenic plants indicated an elevation in the leaf temperature. The thermal status of transformants was similar at both normal and suboptimal water regime. In drought, the transgenic plants used more water during the growing season. The described phenotyping platform provided a comprehensive data set demonstrating the improved physiological condition of the drought stressed transgenic wheat plants in the vegetative growth phase. In soil with reduced water capacity two transgenic genotypes showed higher seed weight per plant than the control non-transgenic one. Limitation of greenhouse-based phenotyping in analysis of yield potential is discussed.     

Distribution and variations of three 1,4-benzoxazin-3-ones in maize induced by the Asian corn borer, Ostrinia furnacalis (Guenée)

Contents of three 1,4-benzoxazin-3-ones in tissue samples from different parts (young leaf, second leaf, old leaf, stem and root) of young maize plants of 4-leaves stage, fed by the third instar larvae of the Asian corn borer, Ostrinia furnacalis (Guenée), were analyzed by high-performance liquid chromatography-mass spectroscopy (HPLC-MS). Samples were taken immediately (set A) or 48 h (set B) after larvae had fed on the second leaf for 48 h. The three 1,4-benzoxazin-3-ones investigated in our experiments were 2,4-dihydroxy-7-methoxy-1,4(2H)-benzoxazin-3-one (DIMBOA), 2,4-dihydroxy-1,4(2H)-benzoxazin-3-one (DIBOA) and 2-hydroxy-7-methoxy-1,4(2H)-benzoxazin-3-one (HMBOA). In samples of set A, the levels of DIMBOA and HMBOA were significantly lifted in the old leaf (L3) and young leaf (L1), respectively, while amounts of these two chemicals in other plant parts were not significantly different between larvae-fed plants and intact plants. Concentrations of DIBOA in each plant part remained unchanged. In samples of set B, no concentration differences for any of these three 1,4-benzoxazin-3-ones between larvae-fed plants and controls were observed in any plant part. The feeding of the Asian corn borer seems to have limited effects on induction of these three 1,4-benzoxazin-3-ones in young maize plants of the variety investigated.     

Effects of Mineral Nutrition on Water Status and Osmotic Adjustment in Spring wheat under Different Moisture Conditions

The Ψw and RWC of the leaves of spring wheat at low level of nutrition were higher than those of the spring wheat leaves at high level during the slow drought period. Therefore, the plants at high nutrition level were more sensitive to drought. High mineral nutrition did not enhance the maximum osmotic adjustment but changed the regulatory process of osmotic adjustment under drought condition. During moderate to nearly severe drought period higher osmotic adjustment was shown in the leaves of spring wheat at high nutrition level. The major solute contributed to osmotic adjustment in the leaves of the spring wheat was soluble sugar, mostly reducing sugar.     

Effects of water deficit on photosynthetic rate and osmotic adjustment in tetraploid wheats

Osmotic adjustment, accumulation of soluble saccharides, and photosynthetic gas exchange were studied in five durum wheat (Triticum turgidum L. var. durum) and one wild emmer wheat (Triticum turgidum L. var. dicoccoïdes) cultivars of contrasting drought tolerance and yield stability. Soil water contents (SWC) were 100, 31, 20, and 12 % of maximum capillary capacity. Under mild water stress (SWC 31 to 20 %), osmotic adjustment capacity and high accumulation of saccharides were found in cv. Cham1, a high yielding and drought tolerant cultivar, and in var. dicoccoïdes, while lowest values were noted in the durum wheat landraces Oued-Zenati and Jennah-Khotifa. Under more severe water stress (SWC 12 %), the cv. Cham1 maintained higher net photosynthetic rate (PN) than other genotypes. The observed changes in the ratio intercellular/ambient CO2 concentration (ci/ca) indicated that under mild and severe water stress, the decrease in PN was mainly due to stomatal and non-stomatal factors, respectively.     

The influence of the mycorrhiza Glomus etunicatum on drought acclimation in safflower and wheat

A study was done to determine the effects of vesicular‐arbuscular mycorrhizal (VAM) colonization on drought acclimation of host plants. Safflower ( Carthamus tinctorius L. cv. S555) and wheat ( Triticum aestivum L. cv. Anza) were grown under environmentally controlled conditions with or without the VAM fungus, Glomus etunicatum Becker and Gerd., and were either acclimated (by pre‐exposing plants to a 10–11 day drought period) or unacclimated to drought. Plants from all treatments were then exposed to drought for 9 days, and plant water status and root water uptake were measured. To minimize interactions between drought and P uptake, growth periods were adjusted so that acclimated and unacclimated plants were similar in size when measurements were made. When wheat was acclimated to drought, osmotic adjustment occurred (leaf solute potentials of leaf tissue rehydrated to full turgor were approximately 0.5 MPa lower in acclimated than unacclimated plants); in safflower, osmotic adjustment was minimal when plants were acclimated. Consequently, acclimated wheat plants were able to tolerate drought better than unacclimated plants, and maintained higher leaf water potentials and relative water contents as soil water was depleted. For both safflower and wheat, acclimated plants had higher water use efficiency, and therefore produced more biomass when water availability was limited, than unacclimated plants. However, mycorrhizal colonization did not affect osmotic adjustment, plant water status, water use efficiency or water uptake in either plant species, and therefore had no effect on drought acclimation or resistance.     

Increase of glycinebetaine synthesis improves drought tolerance in cotton

The tolerance to drought stress of the homozygous transgenic cotton (Gossypium hirsutum L.) plants with enhanced glycinebetaine (GB) accumulation was investigated at three development stages. Among the five transgenic lines investigated, lines 1, 3, 4, and 5 accumulated significantly higher levels of GB than the wild-type (WT) plants either before or after drought stress, and the transgenic plants were more tolerant to drought stress than the wild-type counterparts from young seedlings to flowering plants. Under drought stress conditions, transgenic lines 1, 3, 4, and 5 had higher relative water content, increased photosynthesis, better osmotic adjustment (OA), a lower percentage of ion leakage, and less lipid membrane peroxidation than WT plants. The GB levels in transgenic plants were positively correlated with drought tolerance under water stress. The results suggested that GB may not only protect the integrity of the cell membrane from drought stress damage, but also be involved in OA in transgenic cotton plants. Most importantly, the seedcotton yield of transgenic line 4 was significantly greater than that of WT plants after drought stress, which is of great value in cotton production.